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THE VELIGER # The Veliger 51(1):85–103 (January/April XX, 2008) CMS, Inc., 2008

Trophonella (: ), a New from Antarctic Waters, with the Description of a New Species

M. G. HARASEWYCH Department of Zoology, MRC-163, National Museum of Natural History, Smithsonian Institution, P.O. Box 37012, Washington, DC 20013-7012, USA (e-mail: [email protected])

GUIDO PASTORINO Museo Argentino de Ciencias Natureles, Av. Angel Gallardo 470, 3u piso, lab. 57, C1405DJR Buenos Aires, Argentina (e-mail: [email protected])

Abstract. The new genus Trophonella is described from the outer shelf and upper continental slope of Antarctica and islands within the Antarctic Convergence. Four previously known species that had been attributed to the genus (Trophon scotianus Powell, 1951; T. echinolamellatus Powell, 1951; T. enderbyensis Powell, 1958; and T. eversoni Houart, 1997) are included in Trophonella, as is one new species (Trophonella rugosolamellata) described herein. Trophonella resembles Trophon in gross shell morphology: the members of both genera have large, globose shells, paucispiral protoconchs, prominent axial lamellae, and short siphonal canals. Trophonella differs from Trophon in having shells with evenly rounded whorls that lack a well-defined shoulder; rachidian teeth with distinctive, broadly triangular central cusps, but that lack the marginal cusps of Trophon; characteristic spherical accessory salivary glands; and a circumpapillar fold on the penis that is absent in Trophon. Relationships of the genera Trophon and Trophonella, as well as of the subfamily are reexamined by supplementing the data matrix of Kool (1993b, Table 3) with data for additional taxa. Results support the segregation of Trophonella from Trophon at the generic level. Based on the relationships of the type species of their respective nominotypical genera, Trophoninae is either the sister taxon of a narrowly circumscribed , or both are part of a larger clade. A better resolved phylogeny containing a much broader sampling of the more than 50 genus-level taxa that have been attributed to these two subfamilies will be required in order to delineate more precisely the membership of the clade and to identify its diagnostic synapomorphies.

INTRODUCTION site of assignment of many dissimilar forms that seem, on the basis of the shell and radular morphology of the Although the family Muricidae is best known for its type species of Trophon (T. geversianus), unlikely to temperate and tropical shallow-water representatives, it belong there.’’ The genus Aspella is presently assigned also includes a group of taxa inhabiting polar seas and to the subfamily Muricinae on the basis of radular intervening deeper waters that have generally been morphology (Radwin & D’Attilio, 1976, p. 21). These grouped in the subfamily Trophoninae. Cossmann authors went on to exclude from the subfamily (1903) proposed the subfamily Trophoninae to contain Trophoninae a number of genera (e.g., the genera Trophon Montfort, 1810, and Aspella Dall, 1902; Forreria; Zacatrophon Hertlein & Strong, Mo¨rch, 1877. The genus Trophon was subdivided into 1951) because they ‘‘… had radulae characteristic of the the subgenera Trophon and Bucquoy and family Thaididae.’’ In a phylogenetic study, Kool Dautzenberg, 1882. Under Trophon sensu stricto,he (1993a, fig. 65) showed that, based on the morphology included the sections Trophon, Xanthochorus Fischer, of its type species, the genus Trophon was closely 1884, and Forreria Jouseaume, 1880. Trophonopsis was related to the genera Nucella and Ocenebra,andhe further subdivided into Trophonopsis and suggested that future studies would reveal Trophoninae Fischer, 1884. The genus Aspella was not subdivided. to be polyphyletic. Over the past several decades, there has been general Our continuing studies on the Muricidae represented agreement among researchers that Trophoninae is not in the collections of the United States Antarctic a monophyletic group. As was noted by Radwin and Program (USAP) have revealed that four species D’Attilio (1976, p. 175), Trophoninae ‘‘has been the previously described in the genus Trophon (i.e.,

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Trophon scotianus Powell, 1951; T. echinolamellatus phon geversianus, Ocenebra erinaceus, Trophonella Powell, 1951; T. enderbyensis Powell, 1958; and T. scotiana, Boreotrophon aculeatus,andPaziella pazi. eversoni Houart, 1997) appear to be closely related to Harasewych (1984) was the source of data for T. each other, but they differ substantially from the type geversianus, B. aculeatus,andP. pazi.DataforT. species of the genus Trophon. In this paper, we propose geversianus and O. erinaceus were obtained from Kool the new genus Trophonella to include these species, and (1993a). Data on shell ultrastructure of Boreotrophon we describe an additional new species that we attribute aculeatus and Paziella pazi were newly obtained from to this genus. A review of each of the species included the voucher material from Harasewych’s (1984) study. in this new genus is provided, based on more recently Each of the newly added taxa were scored for the 18 collected specimens. characters and their character states, as defined in Kool We also examine the relationships of the genus (1993b). The data were analyzed using maximum Trophon and the subfamily Trophoninae by reanalyz- parsimony (MP) and neighbor-joining algorithms of ing Kool’s (1993b, table 3) data matrix of primarily PAUP* 4.0 Macintosh Beta Version 10 (Swofford, ocenebrine and rapanine taxa supplemented by the 2002). addition of data for (1) Trophon geversianus (Pallas, 1769), the type species of the type genus of Trophoni- SYSTEMATICS nae; (2) Ocenebra erinaceus (Linnaeus, 1758), the type species of the type genus of Ocenebrinae; (3) Tropho- Class GASTROPODA Cuvier, 1797 nella scotiana (Powell, 1951), the type species of the new Order Wenz, 1938 genus Trophonella; (4) Boreotrophon aculeatus Watson, 1882; and (5) Paziella pazi (Crosse, 1869), the type Family MURICIDAE Rafinesque, 1815 species of the genus Paziella. Trophonella new genus

MATERIALS AND METHODS Type species: Trophon scotianus Powell, 1951. This study is based primarily on specimens collected for Diagnosis: Shell large, fusiform, inflated. Protoconch the United States Antarctic Program (USAP) aboard paucispiral (1.5–2 whorls). Teleoconch (6–7 whorls) the vessels R/V Hero, R/V Eltanin and R/V Professor globose, without distinct shoulder, with tall, conical Sedlacki and housed in the collections of the National spire. Sculpture of low, rounded, spiral cords and axial Museum of Natural History, Smithsonian Institution lamellae that may be broadly flaring to short and (USNM). Additional specimens from several Antarctic rugose. Aperture broadly rounded, with flaring outer expeditions of the Argentine Republic deposited in the lip, and thin inner lip. Siphonal canal short. Operculum Museo Argentino de Ciencias Naturales (MACN) were large, oval, with subterminal nucleus. Rachidian teeth examined, as were specimens at the Zoological Institute with rectangular basal plate; three major cusps, central and Museum, Hamburg (ZMH), and the Senckenberg cusp exceptionally broad, triangular, with small denti- Museum, Frankfurt (SMF), collected by the vessels R/ cle on each side; marginal cusps absent. Accessory V Walther Herwig and R/V Polarstern. All specimens salivary glands small, spherical. Esophageal gland were compared with the holotypes of their respective reduced. Penis with conical papilla surrounded by species, which are housed in The Natural History collar. Museum, London (NHM), the South Australian Museum (SAM), and at USNM. Description: Shell large (to 76 mm), fusiform, often Alcohol-preserved of most species of Troph- broadly ovate, with strongly convex, evenly rounded onella were dissected, and the gross anatomy of the whorls that do not form a distinct shoulder. Proto- anterior part of the alimentary system and pallial conch paucispiral, occasionally with weak, irregular, gonoducts were compared. Radulae were prepared spiral cords. Teleoconch sculpture of low, rounded, according to the method described by Solem (1972) and spiral cords with or without scales, and axial lamellae examined using a Scanning Electron Microscope that range from broadly flaring to short and rugose. (SEM). Terminology for rachidian-tooth morphology Aperture large, suboval, with thick outer lip, and thin follows Kool (1987, fig. 1). Shell ultrastructure data inner lip. Siphonal canal short, scabrous. Shell was obtained from fracture surfaces of shell fragments ultrastructure of three layers, the inner two of removed from the central portion of the lip along the orthogonally oriented crossed-lamellar aragonite, the last shell whorl. outermost layer of calcite. Operculum large, filling In order to determine the generic and subfamilial aperture, corneous, oval, with subterminal nucleus, relationships of this distinctive group of Antarctic thick rim, and large attachment area. muricids, we reanalyzed the data matrix in Kool large, unpigmented. Cephalic tentacles wide, (1993b, table 3), to which we added data for Tro- blunt. Mantle edge smooth, siphon short. Pleuroem-

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bolic proboscis short, broad. Radular ribbon small, 51; Powell, 1960, p. 154; Okutani, 1986, p. 279, pl. 1, thin, extending beyond rear of buccal mass. Rachidian figs. 3–6; Dell, 1990, p. 207, figs. 346–347; Castellanos teeth with subrectangular, weakly recurved basal plate & Landoni, 1993, p. 13, fig. 36; Numanami, 1996, and smooth, broad, marginal surfaces that lack p. 138, figs. 91 a–c. marginal cusps. Central cusp long, exceptionally broad, triangular; flanked by a single, small to nearly obsolete Trophon sp. 1 Hain, 1990, p. 63, pl. 6, fig. 9 a,b; pl. 25, lateral denticle on each side; lateral cusps short, robust, fig. 2. smooth. Lateral teeth of moderate size, basal plate shorter and narrower than in rachidian teeth, with Description: Shell (Figures 1–6) large (to 70 mm), single scythelike cusp along outer edge that is slightly fusiform, thin shelled, chalky. Protoconch (Figures 7– longer than the basal plate. Esophagus short. Salivary 10) tall, increasing in diameter from 450 mmto1.2mm glands very large, ascinous, enveloping esophagus. in 1.5 to 1.75 evenly rounded whorls, surface with very Accessory salivary glands small, spherical, and com- fine, irregular and discontinuous spiral threads. Tran- pletely embedded in corresponding salivary gland. sition to teleoconch marked by broadly flaring lip. Gland of Leiblein compact, with terminal ampula. Teleoconch of up to 6 slightly globose whorls; spire tall, Penis large, wide, with conical papilla rising from a conical, less than 50% of shell length. Suture impressed concavity at its distal end. to abutting. Spire angle 62u–84u. Aperture subpolygo- Etymology: Trophon (Gr., that which feeds) + ella (L., nal to evenly rounded, deflected from the coiling axis suffix added to form the diminutive). by 20u–25u; interior glossy. Siphonal canal moderately short, narrow, slightly deflected dorsally. Siphonal Included species: Trophonella scotiana (Powell, 1951), fasciole scabrous; pseudoumbilical chink nearly closed new combination [Type species]; T. echinolamellata or reduced to a very narrow slit. Outer lip rounded with (Powell, 1951), new combination; T. eversoni (Houart, reflected edges. Columellar lip narrow, adpressed. 1997), new combination; T. enderbyensis (Powell, Axial sculpture of regular, thin strongly developed 1958), new combination; T. rugosolamellata,new lamellae (4–11 on last whorl) that span whorl surface species. from suture to siphon, expanded in shoulder region. Fine, closely spaced growth lines cover whorls and Remarks: Trophonella is similar to Trophon in gross lamellae. Spiral sculpture, of low, rounded, spiral shell morphology, as both have globose shells of cords, broader than interspaces (18–32 on body whorl, comparable size with paucispiral protoconchs, promi- 2–6 on siphonal canal) becoming reduced to obsolete nent axial lamellae, and short siphonal canals. Species on larger specimens. Shell color white to pale orange- of Trophonella can be distinguished most readily by tan. When color present, lighter bands may be evident, their evenly rounded whorls that lack a well-defined one below suture, the other above the juncture to the shoulder. The main differences appear to be anatom- siphonal canal. ical. Trophonella has a circumpapillar fold on the penis Shell composed of three layers (Figure 11). Inner- that is absent in Trophon. The accessory salivary glands most layer, of crossed lamellar aragonite with crystal of Trohonella are globular, while those of Trophon are planes oriented perpendicular to the growing edge, elongate. comprises roughly 2 of shell thickness. Middle layer, The rachidian teeth of Trophonella have distinctively % also aragonitic, arranged with crystal planes parallel to broadly triangular central cusps and lack the marginal growing edge, comprises roughly 36 of shell thick- cusps of Trophon. % ness. Outermost layer is calcitic, comprising the major Trophonella is restricted in geographic distribution to part (62 ) of the shell’s thickness. Operculum (Fig- Antarctica and islands within the Antarctic Conver- % ure 43) oval, with terminal nucleus, outer surface with gence. Records range in depth from 18 m to 474 m regular growth lines, inner surface with broad, thickly (both for T. scotiana), with the majority of specimens glazed rim along posterior edge, and large, round/oval sampled at outer-shelf and upper continental-slope attachment area, often with 2–3 horseshoe-shaped depths (Figures 20–21). In terms of size and biomass, scars. species of Trophonella are the largest muricids occur- ring within the Antarctic Convergence. Animal large, compact. Foot short, broad, with accessory boring organ situated along ventral midline, Trophonella scotiana (Powell, 1951), opening via ventral pedal gland in females. Tentacles new combination large, broad, closely situated, with large black eyes. Mantle edge smooth, osphradium (Figure 48, os) (Figures 1–13, 20) small, slightly asymmetrical, with 45–55 leaflets per side, less than 50% the length of the ctenidium Synonymy: Trophon scotianus Powell, 1951, p. 153, pl. (Figure 48, ct). Proboscis large, muscular, cylindrical, 9, figs. 48–49, M88; Carcelles, 1953, p. 189, pl. 2, fig. pleurombolic. Radula (Figures 12–13) short (radular

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Figures 1–11. Trophonella scotiana (Powell, 1951). 1. Apertural, 2. lateral, and 3. dorsal views of a medium-sized specimen with few axial lamellae, USNM 897411, off South Georgia Island, 54u539S, 35u499W, in 127–140 m. 4. Apertural view of large specimen with well-developed lamellae, USNM 901644, Off Borchgrevink Coast, Victoria Land, Antarctica, 74u019S, 178u539E, in 256– 258 m. 5. Apertural and 6. dorsal views of average specimen, USNM 897560, off South Georgia Island, 54u399S, 37u229W, in 140– 150 m. 7–8. Lateral, and 9. apical views of the protoconch. 10. Detail of protoconch sculpture. USNM 901645, W of Graham Land,

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length roughly 30% of aperture length), narrow 1975; USNM 887835, 1 L, off South Georgia Island, (roughly 550 mm wide), extending beyond rear of 54u019180S, 36u509420W, 108–119 m, R/V Islas Orca- retracted proboscis. Rachidian tooth subrectangular, das, cr. 575, stn. 24, 17 May 1975; USNM 896099, 4 D, basal plate moderately concave anteriorly, with short, off South Georgia Island, 53u519S, 37u389W, in 97– smooth marginal area. Central cusp large, broad, 101 m, R/V Eltanin cr. 22, stn. 1535, 7 Feb 1966; lateral cusps shorter, narrower, with single denticle USNM 896982, 1 L, off South Georgia Island, 53u579S, between central and lateral cusps (Figure 13, d) often 36u069W, 151–158 m, R/V Professor Siedlecki, cr. 601, almost obsolete, but present, even in juvenile speci- stn. 97, 13 Dec 1986; USNM 897411, 2 L, off South mens. Lateral teeth L-shaped, with single cusp along Georgia Island, 54u539S, 35u499W, in 127–140 m, R/V outer edge roughly equal in length to width of basal Professor Siedlecki, cr. 601, stn. 71, 9 Dec 1986; USNM plate. Juvenile specimens have proportionally larger 897416, 1 L, off South Georgia Island, 55u249S, denticles, and thinner cusps on rachidian and lateral 35u229W, 207–218 m, R/V Professor Siedlecki, cr. teeth. Esophagus very thin. Valve of Leiblein (Fig- 601, stn. 60, 8 Dec 1986; USNM 897422, 1 L, off ure 49, vL) ovate. Salivary glands (Figure 49, sg) very South Georgia Island, 55u089S, 35u069W, 145–153 m, large, envelop the valve of Leiblein and the small, R/V Professor Siedlecki, cr. 601, stn. 66, 9 Dec 1986; nearly spherical accessory salivary glands, joining USNM 897427, 2 L, off South Georgia Island, 55u019S, esophagus via two thick ducts anterior to the valve of 35u279W, 117–122 m, R/V Professor Siedlecki, cr. 601, Leiblein. Mid-esophagus very thin, lacks conspicuous stn. 67, 9 Dec 1986; USNM 897433, 2 L, off South esophageal gland, passes under gland of Leiblein. Georgia Island, 55u059S, 35u239W, 116–121 m, R/V Gland of Leiblein large, compact, with blind duct Professor Siedlecki, cr. 601, stn. 68, 9 Dec 1986; USNM terminating posteriorly in large ampulla. Stomach 897434, 5 L, off South Georgia Island, 54u319S, (Figures 50, 51, sto) large, lining the anterior and 38u119W, 180–190 m, R/V Professor Siedlecki, cr. dorsal surface of the digestive gland, into which it 601, stn. 36, 4 Dec 1986; USNM 897444, 1 L, off opens via ducts (Figure 51, dd). Intestine (Figures 50, South Georgia Island, 53u559S, 37u089W, 104–122 m, 51, i) broad. Penis (Figures 45–47, p) large (. 4 3 R/V Professor Siedlecki, cr. 601, stn. 109, 14 Dec 1986; tentacle length), broad, dorsoventrally flattened, elon- USNM 897459, 1 L, off South Georgia Island, 53u529S, gate, with laterally situated sperm duct, dorsal blood 37u339W, 104–112 m, R/V Professor Siedlecki, cr. 601, sinus, and conical papilla (Figures 46, 47, 52, pp) stn. 111, 15 Dec 1986; USNM 897475, 3 L, off South surrounded by a collar. Pallial oviduct broad. Albumin Georgia Island, 53u529S, 38u209W, 113–120 m, R/V gland thin, capsule gland (Figure 53, cg) short, globose. Professor Siedlecki, cr. 601, stn. 120, 16 Dec 1986; Bursa copulatrix (Figure 53, bc) onion-shaped, with USNM 897490, 1 L, off South Georgia Island, 54u119S, distal female opening (Figure 53, fo). Rectum (Fig- 37u539W, 113–118 m, R/V Professor Siedlecki, cr. 601, ure 53, r) overlays pallial oviduct; rectal gland (Fig- stn. 40, 5 Dec 1986; USNM 897521, 2 L, off South ure 53, rg) lines anterior dorsal surface of rectum; anus Georgia Island, 54u059S, 38u259W, 197–207 m, R/V (Figure 53, a) adjacent to female opening. Egg capsules Professor Siedlecki, cr. 601, stn. 24, 3 Dec 1986; USNM of T. scotiana lenticular, subcircular in outline, 897560, 1 L, off South Georgia Island, 54u399S, attached to substrate (Hain, 1990, pl. 6, fig. 9b, 37u229W, 140–150 m, R/V Professor Siedlecki, cr. Trophon sp. 1). Each capsule is about 19 mm in 601, stn. 46, 6 Dec 1986; USNM 897564, 5 L, off diameter, contains 140 eggs that hatch as crawling South Georgia Island, 54u519S, 35u389W, in 84–103 m, juveniles in 24–25 months (Hain & Arnaud, 1992, R/V Professor Siedlecki, cr. 601, stn. 72, 9 Dec 1986; p. 307, table 3). USNM 901642, 1 D, Ross Sea, Moubray Pennell Bank, Victoria Land, Antarctica, 73u229S, 177u379E, in 465– Type locality: Off South Georgia Island, 53u559S, 474 m, R/V Eltanin, cr. 27, stn. 1933, 30 Jan 1967; 38u019W, in 107 m (Figure 20, ). USNM 901644, 1 L, Ross Sea, Moubray Pennell Bank, Type material: Holotype, NHM 1961542. Victoria Land, Antarctica, 74u019S, 178u539E, in 256– 258 m, R/V Eltanin, cr. 32, stn. 2018, 14 Jan 1968; Material examined: D indicates empty shells, L USNM 901645, 1 L, W of Graham Land, Palmer indicates live-collected specimens. Peninsula, Antarctica, 66u21.79S, 66u479W, in 70– NHM 1961542, Holotype; USNM 887834, 1 L, off 106 m, R/V Hero, cr. 731, stn. 1861, 1 Mar 1973; South Georgia Island, 53u509360S, 36u189360W, 185– MACN-In 18991, 1 D, Schlieper Bay, South Georgia 205 m, R/V Islas Orcadas, cr. 575, stn. 30, 19 May Island, in 18 m; MACN-In 18990, 1 L, Cumberland

r

Palmer Peninsula, Antarctica, 66u21.79S, 66u479W, in 70–106 m. Scale bars 5 400 mm. 11. Shell ultrastructure. Fracture surface parallel to growing edge. USNM 887835, off South Georgia Island, 54u019180S, 36u509420W, 108–119 m.

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Figures 12–19. Radulae of species of Trophonella. 12. Dorsal and 13. right lateral views of the radular ribbon of T. scotiana, specimen in Figure 4. 14. Dorsal and 15. right lateral views of the radular ribbon of T. echinolamellata, USNM 896041, 2L, NW of Brabant Island, Palmer Archipelago, Antarctic Peninsula, 63u519S, 62u389W, in 128–165 m. 16. Dorsal and 17. right lateral views of the radula of T. enderbyensis, specimen in Figures 38–40. 18. Dorsal and 19. right lateral views of the radula of the holotype of T. rugosolamellata n. sp. ZMH 2777.

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Figure 20. Geographic and bathymetric distributions of Trophonella scotiana, T. rugosolamellata n. sp., and the only locality reported for T. eversoni.

Bay, South Georgia Island, in 36 m; ZMH w/n, 1 D, R/V Walther Herwig, cr. 68, stn. 64; ZMH w/n, 1 L, off off South Georgia Island, 54u13.29S, 37u49.69W, in South Georgia Island, 54u489S, 35u23.69W, in 215 m, 122 m, R/V Walther Herwig, cr. 68, stn. 15; ZMH w/n, R/V Walther Herwig, cr. 68, stn. 70; ZMH w/n, 4 L, off 1 L, off South Georgia Island, 53u55.99S, 38u29.69W, in South Georgia Island, 55u15.29S, 34u46.89W, in 240 m, 145 m, R/V Walther Herwig, cr. 68, stn. 19; ZMH w/n, R/V Walther Herwig, cr. 68, stn. 78; ZMH w/n, 2 L, off 3 L, 1 D, off South Georgia Island, 53u39.59S, South Georgia Island, 55u0.379S, 35u03.89W, in 124 m, 37u8.99W, in 155 m, R/V Walther Herwig, cr. 68, stn. R/V Walther Herwig, cr. 68, stn. 79; SMF w/n, 1 D, off 30; ZMH w/n, 1 D, off South Georgia Island, the Princess Martha Coast, Queen Maud Land, 53u47.39S, 37u12.29W, in 141 m, R/V Walther Herwig, Antarctica, 70u299S, 8u079W, in 270–303 m. cr. 68, stn. 31; ZMH w/n, 3 L, off South Georgia Island, 53u59.89S, 36u56.29W, in 150 m, R/V Walther Literature records: PS ANT V/3 St. 593, 73u559S, Herwig, cr. 68, stn. 35; ZMH w/n, 1 L, off South 23u389W, in 330 m (Hain 1990); JARE-25, St. A, Georgia Island, 53u43.39S, 36u269W, in 230 m, R/V 70u149S, 24u23.99E, Breid Bay, in 310 m (Okutani 1986; Walther Herwig, cr. 68, stn. 40; ZMH w/n, 1 L, off Numanami 1996). South Georgia Island, 53u59.19S, 36u20.69W, in 198 m, R/V Walther Herwig, cr. 68, stn. 44; ZMH w/n, 1 L, off Distribution: Off South Georgia Island, the Antarctic South Georgia Island, 53u52.99S, 36u4.29W, in 235 m, Peninsula, and the Ross Sea, the Weddell Sea, and R/V Walther Herwig, cr. 68, stn. 55; ZMH w/n, 4 L, off Breid Bay, Antarctica, at depths of 18–474 m. See South Georgia Island, 54u32.69S, 35u58.19W, in 155 m, Figure 20.

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Remarks: This distinctive taxon, the largest species of thickness), consisting of crossed lamellar aragonite with Trophonella in terms of biomass, had, until recently, crystal planes oriented perpendicular to growing edge been known from less than a dozen specimens (Dell, of the shell. Middle layer of crossed lamellar aragonite, 1990:208; Numanami, 1996:138). A survey of the with crystal planes colabrally oriented, accounts for collections of Antarctic mollusks at USNM, SMF approximately 30% of shell thickness. Outermost layer and ZMH, as well as a search of the literature, has thickest (, 68% of shell thickness), calcitic. Operculum uncovered more than 60 specimens and/or records, (Figure 42) D-shaped, with straight adaxial margin, most from the vicinity of South Georgia Island and the terminal nucleus, fine growth lines along external Weddell Sea. Dell (1990:208) and Numanami surface, and broad, glazed rim along inner surface. (1996:140) both noted variation in shell morphology Attachment area without scars. of this species, with the degree of whorl inflation and The animal is similar to that of T. scotiana in most the number of axial lamellae per whorl both increased respects. The radula (Figures 14, 15) is slightly longer in large specimens. (radular length , 0.45 aperture length) and has proportionally broader rachidian teeth with more Trophonella echinolamellata (Powell, 1951) pronounced denticles. new combination Type locality: Off Cape Bowles, Clarence Island, (Figures 14, 15, and 21–28) Antarctica, 61u259S, 53u469W, in 342 m. (Figure 21, ,).

Synonyms: Trophon echinolamellatus Powell, 1951:152, Type material: Holotype and one paratype, NHM pl. 9, figs. 44, 45. Trophon equinolamellatus [sic] 1961541. Pastorino, 2002:fig. 22. Material examined: D indicates empty shells, L Description: Shell (Figures 22–24) large (to 69.8 mm), indicates live collected specimens. fusiform, inflated, thin shelled, with scabrous surface. NHM 1961541, Holotype and one paratype; USNM Protoconch (Figures 25–27) tall, increasing in diameter 638874, 1 L, off Victor Hugo Island, west coast of from 300 to 920 mm in 1.75 to 2 evenly rounded whorls, Palmer Peninsula, Antarctica, 65u089S, 66u049W, in surface smooth, frequently pitted. Transition to tele- 135 m, stn. ED28, 22 Mar 1959; USNM 678397, 3 L, oconch distinguished by weak flare in lip. Teleoconch off Palmer Peninsula, Antarctica, 60u489S, 44u13.5W, of up to 7 evenly rounded whorls; spire tall, conical, in 188 m, R/V Eastwind, stn. EW66-028, 11 Feb 1966; slightly less than half the shell length. Suture distinctly USNM 846180, 1 D, east of Victor Hugo Island, west impressed. Spire angle 66–72u. Aperture broadly ovate, coast of Palmer Peninsula, Antarctica, 65u049S, deflected from the coiling axis by about 25u, interior 65u539W, in 150 m, R/V Polar Duke, 8 Sept 1985; with glazed, lustrous white layer that does not extend to USNM 870316, 1 L, NW of Brabant Island, Palmer the end of the flared, rounded, outer lip, nor to the edge Archipelago, Antarctic Peninsula, 63u519S, 62u389W, of thick, adpressed, columellar lip. Demarcation 128–165 m, R/V Eltanin, cr. 6, stn. 439, 9 Jan 1963; between apertural glaze and apertural lip very pro- USNM 870593, 2 L, NE of Joinville Island, Antarctic nounced, especially in pigmented shells. Siphonal canal Peninsula, 62u419S, 54u439W, 210–220 m, R/V Eltanin, short, narrow, slightly deflected dorsally. Siphonal cr. 12, stn. 1003, 15 Mar 1964; USNM 881728, 1 L, off fasciole scabrous; pseudoumbilical chink pronounced, Visokoi Island, South Sandwich Islands, 56u429180S, constricted to varying degrees by thick columellar lip. 27u009240W, 93–121 m, R/V Islas Orcadas, cr. 575, stn. Outer lip rounded, weakly reflected. Columellar lip 61, 30 May 1975; USNM 881906, 1 L, D’Urville Island, thick, overlaying sculpture of previous whorl. Axial Bransfield Strait, Antarctic Peninsula, 62u399S, sculpture of numerous (14–16 on first teleoconch 56u109W, 426–311 m, R/V Eltanin, cr. 6, stn. 418, 2 whorl, 42–50 on last whorl), extremely short lamellae Jan 1963; USNM 881907, 1 L, D’Urville Island, that intersect with closely spaced spiral cords that are Bransfield Strait, Antarctic Peninsula, 62u399S, as broad as interspaces (16–20 on body whorl, 2–4 on 56u109W, 426–311 m, R/V Eltanin, cr. 6, stn. 418, 2 siphonal canal) to produce a scaly surface. On the final Jan 1963; USNM881939, 1 L 2 D, off South Georgia whorl of very large specimens, several lamellae may be Island, 54u419S, 38u389W, 220–320 m, R/V Eltanin, cr. close together and thickened, giving the appearance of 9, stn. 671, 23 Aug 1963; USNM 896041, 2 L, NW of varices that may be spaced 0.25 to 0.5 whorl apart. Brabant Island, Palmer Archipelago, Antarctic Penin- Shell white, or, more commonly pale orange-cinnamon. sula, 63u519S, 62u389W, in 128–165 m, R/V Eltanin, cr. When color is present, lighter bands may be evident, 6, stn. 439, 9 Jan 1963; USNM 896056, 2 L, E of South one below the suture, the other above the juncture to Orkney Islands, 60u519S, 42u579W, in 155 m, R/V the siphonal canal. Eltanin, cr. 12, stn. 1083, 14 Apr 1964; USNM 901636, The shell is composed of three layers (Figure 28). 2 D, off Zavodovski Island, South Sandwich Islands, Innermost layer very thin (approximately 2% of shell 56u28.89S, 27u24.69W, in 161–210 m, R/V Islas Orca-

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Figure 21. Geographic and bathymetric distributions of Trophonella echinolamellatus and T. enderbyensis.

das, cr. 575, stn. 70, 2 Jun 1975; USNM 901638, 1 L, of Laurie Island, South Orkney Islands, Antarctica, Gibbs Island, Bransfield Strait, South Shetland Islands, 60u41.69S, 43u57.19W 290 m, R/V Walther Herwig, cr. Antarctica, 61u259S, 56u309W, in 300 m, R/V Eltanin, 68, stn. 88; ZMH w/n, 1 D, NNE of Laurie Island, > cr. 12, stn. 998, 14 Mar 1964; USNM 901639, 1 L, Low South Orkney Islands, Antarctica, 60u34.99S, Island, South Shetland Islands, Antarctica, 63u269S, 44u17.39W 240 m, R/V Walther Herwig, cr. 68, stn. 62u159W, in 119–124 m, R/V Hero, cr. 691, stn. 26, 10 89; ZMH w/n, 4 L, 2 D, SE of Laurie Island, South ? Feb 1969; USNM 901640, 1 L, Aspland Island, South Orkney Islands, Antarctica, 60u51.39S, 44u129W 178 m, Shetland Islands, Antarctica, 61u179S, 56u269W, in R/V Walther Herwig, cr. 68, stn. 90; ZMH w/n,3L, @ 421–462 m, R/V Polarstern, cr. 2, stn. 22, 21 Nov 1 D, WNW of Coronation Island, South Orkney 1996; USNM 901641, 4 L, Elephant Island, South Islands, Antarctica, 60u259S, 46u25.79W 150 m, R/V Shetland Islands, Antarctica, 60u539S, 55u329W, in Walther Herwig, cr. 68, stn. 118; ZMH w/n, 1L, NW of A 178–120 m, R/V Polarstern, cr. 2, stn. 79, 9 Dec 1996; Aspland Island, South Shetland Islands, Antarctica, col. P. Arnaud, 200–220 m, South Shetland Islands, 61u21.89S, 56u0.69W 368 m, R/V Walther Herwig, cr. ; Antarctica; ZMH w/n, 1 L, NW of Elephant Island, 68, stn. 138; ZMH w/n, 1 L, 3 D, NW of Aspland B South Shetland Islands, Antarctica, 60u569S, Island, South Shetland Islands, Antarctica, 61u19.89S, 55u32.99W 81 m, R/V Walther Herwig, cr. 68, stn. 68; 56u9.99W 328 m, R/V Walther Herwig, cr. 68, stn. 139; < ZMH w/n, 2 L, E of Laurie Island, South Orkney ZMH w/n, 1 L, 1 D, NW of Aspland Island, South Islands, Antarctica, 60u48.99S, 43u349W 257 m, R/V Shetland Islands, Antarctica, 61u12.59S, 56u23.49W = Walther Herwig, cr. 68, stn. 87; ZMH w/n,2L2D,E 460 m, R/V Walther Herwig, cr. 68, stn. 141; ZMH

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Figures 22–28. Trophonella echinolamellata (Powell, 1951). 22. Apertural, 23. lateral, and 24. dorsal views of a large specimen, USNM 901636, Off Zavodovski Island, South Sandwich Islands, 56u28.89S, 27u24.69W; 161–210 m. 25. Lateral, 26. abapertural, and 27. apical views of the protoconch, USNM 870593, NE of Joinville Island, Antarctic Peninsula, 62u419S, 54u439W, 210–220 m. 28. Shell ultrastructure. Fracture surface parallel to growing edge. USNM 901638, off Gibbs Island, Bransfield Strait, South Shetland Islands, Antarctica, 61u259S, 56u309W, in 300 m.

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C w/n, 2L, W of Elephant Island, South Shetland Islands, regular growth lines, inner surface with broad, thickly Antarctica, 61u12.79S, 55u56.49W 134 m, R/V Walther glazed rim along posterior edge. EX Herwig, cr. 68, stn. 148; ZMH w/n, 2 L, W of Elephant A single, male animal was examined and found to be Island, South Shetland Islands, Antarctica, 61u89S, anatomically similar to T. scotiana. The radula 55u56.29W 125 m, R/V Walther Herwig, cr. 68, stn. (Figures 16–17) is similar to that of T. scotiana but EO 150; ZMH w/n, 1 L 1 D, NW of Elephant Island, South has more prominent denticles between the central and Shetland Islands, Antarctica, 60u53.99S, 55u30.49W lateral cusps of the rachidian. 135 m, R/V Walther Herwig, cr. 68, stn. 159. Type locality: (Figure 21:%), off Enderby Land, Literature records: Only the type locality has been Antarctica, 65u489S, 71u249E in 193 m. [British, Aus- reported in the literature previously. tralian, and New Zealand Antarctic Research Expedi- tion, St. 41, January 24/25 1930]. Distribution: Known from Antarctic Peninsula, South Shetland Islands, South Georgia Island, South Sand- Type material: Holotype, SAM D15497. wich Islands and South Orkneys Islands, at depths Material examined: Holotype; SMF w/n, 1 D, Off the ranging from 93 m to 460 m (Figure 21). Princess Martha Coast, Queen Maud Land, Antarc- tica, 70u309S, 8u49W, 261–263 m; SMF w/n, 1 L, off the Remarks: The characteristic ornamentation of the shell, Princess Martha Coast, Queen Maud Land, Antarc- consisting of sharply raised spiral cords overlain by closely tica,71u239S, 13u589W, 293–357 m. spaced lamellae to produce a scabrous surface, distinguish- es T. echinolamellata from all other known species of Literature records: None, apart from the original Trophonella. Trophonella echinolamellata and T. scotiana description. have similar bathymetric ranges, and they overlap in portions of their geographical ranges. However, T. Distribution: Presently known from three specimens, echinolamellata appears to be restricted to the Scotia one from type locality (off Enderby Land) and two tectonic plate, based on the material thus far available. from off Queen Maud Land, Antarctica. Trophonella enderbyensis (Powell, 1958), Remarks: In his description of Trophon enderbyensis, new combination Powell (1958:198) noted that this species resembled T. scotianus, which, at the time, was known only from (Figures 21, 29–31) South Georgia Island. The specimens presently avail- able indicate that the geographic and bathymetric ranges of these two species overlap. Both Dell Synonymy: Trophon enderbyensis Powell, 1958:197, pl. (1990:208) and Numanami (1996:140) commented that 3, fig. 1. larger specimens of T. scotiana tend to have an increased Description: Shell (Figures 29–31) large (to 70 mm), number of axial lamellae and a more inflated final whorl fusiform, moderately thin-shelled, with regular spiral of the shell. The adult holotype of Trophonella sculpture and widely spaced lamellose varices. Proto- enderbyensis is comparable in size to the largest conch and transition to teleoconch unknown. Tele- specimens of T. scotiana, yet it has far fewer, shorter, oconch with 7 or more evenly convex whorls; sutural axial lamellae and a narrower, more fusiform shell. The shelf obsolete; spire tall, narrow, conical, approximate- intermediate denticle on the rachidian teeth of T. ly 50% of shell length; suture abutting; spire angle endebryensis is pronounced, but it is reduced or absent in T. scotiana. As more material becomes available, about 55u. Aperture broadly ovate, deflected from the especially from off Enderby Land and Wilkes Land, the coiling axis by 25u, interior glossy white. Siphonal canal short; siphonal fasciole conspicuous, with recurved, status of T. enderbyensis may require reevaluation. scabrous processes; pseudoumbilical chink reduced to a Trophonella eversoni (Houart, 1997) narrow slit. Outer lip rounded, reflected; columellar lip new combination thin. Axial sculpture of low, generally distantly spaced thin lamellae covering entire whorl, obsolete on early (Figures 32–34) whorls, approximately 3 per whorl except for the final whorl, where 3 lamellae are closely adjacent in the Synonymy: Trophon eversoni Houart, 1997:9, figs. 1–2, holotype. Spiral sculpture of low, rounded cords (3 or 4 4, 6. on early whorls, 20 on last whorl), about as wide as interspaces, which may have 1–4 fine spiral threads Description: Shell (Figures 32–34) large (to 75.8 mm), between adjacent cords. Shell whitish. Shell composed fusiform, of medium thickness, strongly sculptured of three layers, as in T. scotiana. Operculum (Fig- with spiral cords and axial lamellae. Protoconch and ure 44) oval, with terminal nucleus; outer surface with transition to teleoconch unknown. Teleoconch with 6

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Figures 29–31. Trophonella enderbyensis (Powell, 1958). 29. Apertural, 30. lateral, and 31. dorsal views of the holotype, SAM D15497, off Enderby Land, Antarctica, 65u489S, 71u249E in 193 m. 32–34. Trophonella eversoni (Houart, 1997). 32. Apertural, 33. lateral, and 34. dorsal views of the holotype USNM 880177, South Atlantic Ocean, ? Antarctica, ? Kerguelen Is. 35–37. Trophonella rugosolamellata new species. 35. Apertural, 36. lateral, and 37. dorsal views of the holotype, ZMH 2777, off South Orkney Islands, 60u52.29S, 44u28.49W, in 205 m.

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Figures 38–46. Trophonella enderbyensis (Powell, 1958). 38. Apertural, 39. lateral, and 40. dorsal views of a young specimen, SMF w/n, off the Princess Martha Coast, Queen Maud Land, Antarctica, 71u239S, 13u589W, in 293–357 m. 41. T. rugosolamellata new species, operculum of holotype, Figures 35–37. 42. T. echinolamellata (Powell, 1951), operculum of specimen in Figures 22–24. 43. T. scotiana (Powell, 1951), operculum of specimen in Figure 4. 44. T. enderbyensis (Powell, 1958) operculum of specimen in Figures 38–40. 45–46. Critical-point dried penis of T. scotiana (Powell, 1951). 45. Dorsal and left lateral views of penis. 46. Detail of terminal papilla.

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Figures 47–53. Trophonella scotiana (Powell, 1951). 47. Anterior portion of male specimen, mantle reflected. 48. Ctenidium and osphradium. 49. Anterior alimentary system. 50. Dorsal and 51. right lateral views of stomach. 52. Distal tip of penis. 53. Pallial oviduct. Abbreviations for these figures: a, anus; asg, accessory salivary gland; bc, bursa copulatrix; cg, capsule gland; ct, ctenidium; dd, ducts to digestive diverticula; e, esophagus; fo, female opening; i, intestine; mo, mouth; os, osphradium; p, penis; pp, penial papilla; r, rectum; rg, rectal gland; s, siphon; sg, salivary gland; sto, stomach; vl 5 valve of Leiblein.

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or more globose whorls. Sutural shelf present in last Trophonella rugosolamellata new species whorls. Spire conical, about 33% of total shell length. Suture abutting. Spire angle about 56u–59u. Aperture (Figures 20, 35–37) large, suboval, deflected from the coiling axis by 27u; interior glossy white; siphonal canal moderately short, Description: Shell (Figures 35–37) large (to 76 mm), slightly inclined; siphonal fasciole scaly. Outer lip fusiform, solid, chalky, with strong spiral cords and rounded with reflected edges; columellar lip thin, thick axial lamellae that are furrowed and reflected at tightly adpressed, pseudoumbilical chink absent. Axial the shoulder. Protoconch unknown. Teleoconch of at sculpture of pronounced, regularly spaced, thin lamel- least six narrowly convex whorls; spire 33% of total lae, 12–14 on last whorl, running along whorl surface shell length. Suture abutting. Spire angle about 59u. from adapical suture to the siphonal fasciole. Spiral Aperture large, oval, deflected from the coiling axis by sculpture of low, rounded, spiral cords: 3, 8, and 24 on about 22u, interior glossy white. Siphonal canal the first, second, and last whorls, respectively. Cords moderately long and narrow, slightly inclined, siphonal most pronounced along shell periphery, reduced near fasciole strongly scabrous, pseudoumbilical chink suture and siphonal canal. Cords cover the lamellae but closed. Outer lip rounded with reflected edge. Colu- became obsolete along their edges. Shell opaque white. mellar lip broad, with thick, expanded callus. Axial Regular growth lines covering whorls and lamellae. lamellae thick, more or less evenly spaced (11 on last Operculum, radula, and anatomy unknown. whorl, 8 on the penultimate whorl), running along whorl surface from suture to the siphonal fasciole, Type locality: (Figure 20:¤), Antarctica, 185 m. narrowest at midwhorl, most pronounced near and on the siphon and especially at a position corresponding to Material examined: Holotype, USNM 880177. the shoulder, where they are enlarged, furrowed and Literature records: Kerguelen Islands (Paratype 2); strongly reflected. Regular growth lines evident on the submarine bank ‘‘South’’ off the Kerguelen Islands, in whorls and lamellae. Spiral sculpture of rounded, ‘‘approximately’’ 250 m (Houart, 1997). closely spaced cords (3 or 4 on early whorls, 20 on last whorl) that extend onto the outer surfaces of the Distribution: This species is based on a holotype lamellae. Cords most pronounced from siphonal canal collected by a Russian trawler ‘‘in the Antarctic,’’ from to region corresponding to the shoulder. Shell color a depth of 185 m. Paratype 2 is reported, with some whitish. Shell composed of three layers, as in T. certainty, to have been taken off the Kerguelen Islands. scotiana. Operculum (Figure 41) subpolygonal, thick, Houart (1997:9) mentions two additional specimens brownish, with terminal nucleus. Growth lines on from a submarine bank south of the Kerguelen Islands, external surface; broad, glazed rim covering nucleus; at depths of 250 m. attachment area smaller. Animal large. Foot short, broadly rectangular. Remarks: Trophonella eversoni is known from five Tentacles very wide, blunt, with small black eyes; specimens, four of which are in private collections. The mantle edge smooth, siphon very short. Osphradium geographic and bathymetric distributions of this species small, asymmetrical, about 50% ctenidium length. are apocryphal, and they have yet to be confirmed by Pleurembolic proboscis short and broad. Radular subsequent collections. Powell (1957:113) reported the ribbon (Figures 18, 19) short (radular length , 0.44 occurrence of five species of Trophon in the Kerguelen aperture length), thin, extending beyond rear of buccal Islands, although all but two of the stations sampled by mass. Rachidian tooth subrectangular, and basal plate the British, Australian, and New Zealand Antarctic weakly concave anteriorly. Central cusp broadly Research Expedition were from substantially shallower triangular, flanked by short stout denticles. Outer depths. Houart (1997:11) noted that T. albolabratus cusps shorter and narrower, slightly outwardly direct- Smith, 1875, one of the Kerguelen Island species, ed. Marginal cusps absent, marginal area smooth. differs from T. eversoni in having ‘‘low, thin, and more Lateral teeth L-shaped, narrower than rachidian teeth. numerous axial lamellae, stronger spiral cords, and a Esophagus short. Salivary glands large, ascinous, shorter siphonal canal. He distinguished T. eversoni whitish, with ducts entering esophagus just anterior from T. scotiana on the basis of the former having more to the wide valve of Leiblein. Accessory salivary glands numerous spiral cords, more numerous and abapertu- small, spherical, located in the anterior part of the rally sloping axial lamellae that are not as strongly buccal mass, totally embedded in salivary glands. produced adapically. As with T. enderbyensis,addi- Gland of Leiblein compact, without lobules, brownish tional collections from off Enderby Land, Wilkes in color, overlaying esophagus. Penis similar to that of Land, and the islands off their coasts will be required T. scotiana; large, wide, flat, . 4 times tentacle length; to assess more accurately the distribution and relation- with terminal, conical papilla surrounded by narrow ships of T. eversoni. collar.

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Figure 54. Strict consensus maximum parsimony [MP] and neighbor-joining [NJ] trees depicting the relationships of genera based on data derived from their type species. Data from Kool (1993b:Table 3) have been supplemented by data for Ocenebra erinaceus, Trophon geversianus, Trophonella scotiana, Boreotrophon aculeatus,andPaziella pazi.

Type locality: S of Laurie Island, South Orkney Islands, Etymology: Rugosolamellata from Latin rugosus and 60u52.29S, 44u28.49W, in 205 m. Latin lamellata (with lamellae, or thin plates) referes to the extension of the spiral cords onto the axial lamellae. Type material: Holotype, ZMH 2777. Remarks: Trophonella rugosolamellata most closely Material examined: This species is presently known resembles T. eversoni, but it can be distinguished by only from the holotype. its thicker shell, with slightly fewer lamellae and fewer, coarser spiral cords. The axial lamellae of T. rugoso- Distribution: Known only from the type locality, lamellata are distinctive in that they are reflected along collected by the R/V Walther Herwig cr. 68, stn. 91 the outer edges, especially at a position corresponding (Figure 20). to the shoulder, where they are enlarged, furrowed, and

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Table 1 Taxa, characters, and character states used in an analysis of the relationships of Trophoninae. The data matrix is that of Kool (1993b:table 3), to which have been added the taxa Trophon geversianus, Ocenebra erinaceus, Trophonella scotiana, Boreotrophon aculeatus,andPaziella pazi. See Kool (1993b) for detailed descriptions of characters and character states.

Character 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 Taxa from Kool (1993b): Muricanthus aaaaaaaaaa a aaaa a a a Forreria bbbababaaa b aaab ? b c Nucella bbbababaaa b bcab b b c Haustrum b?cabaaaaa b bbae b a b Morula aaaacbabbb c cdbc a a e Cronia caaacbabbb d cdbb a a f Rapana aacadaaabc d edbb a a f Cymia ??caeababa ? ddbb a a d Stramonita aabaeaaabc d edbb a a g Concholepas aabaeaaabc d edbb a a g Dicathais cabaeaabbc ? edbc a a g Vasula ???beaabbcd edbc ? a j Vexilla daaafacbbc d fdbd a a ? Nassa aaaaeaabbc d fdbc a a ? Pinaxia aaaaeaabbc d fdbc a a h Drupa ???aeaabbcd edbda a h Plicopurpura aa?aeaabbc d edbc a a ? Thais ?acbeaabbc d edbc a a j Purpura ?acbeaabbc d edbc a a h Mancinella ??cbeaabbc ? edbc a a i Neorapana ?acbeaabbc d edbc a a j Tribulus ?acbe?ab?? ? ed?? ? a j Acanthina bb?ab?b?a? ? b??? ? ? ? Trochia bbba?????? ? ???? ? ? ? Newly added taxa: Ocenebra erinaceus bbbaa?baaa b bdab b b c Trophon geversianus babaa?aaaa b bcac b b c Trophonella scotiana dabaa?aaaa b baac b a a Boreotrophon aculeatus dacaa?aaaa b baac b a a Paziella pazi bacaa?aaaa a baac b a a

strongly reflected to produce a tapering structure shown in Figure 54 (MP). Figure 54 (NJ) shows the similar to the open shoulder ‘‘spine’’ of Pterochelus neighbor joining tree based on the same data. Jousseaume, 1880. The spiral cords of T. eversoni are Ocenebra erinaceus, the type species of the type genus more numerous and thinner, and they are not as of Ocenebrinae, emerges in an unresolved pentatomy pronounced along the siphonal canal or on the axial with the genera Nucella, Trochia, Forreria,and lamellae as in T. rugosolamellata. Trophonella ender- Acanthina in the strict-consensus tree and as sister byensis has a higher spire, fewer, smaller, and more taxon to these genera in the neighbor-joining tree. widely spaced lamellae, and fainter spiral cords than T. Trophon geversianus, the type species of the type genus rugosolamellata. Trophoninae, emerges as the sister group to this clade in both the MP analysis and the NJ analysis. While the NJ tree shows Haustrum to be the sister taxon to this PHYLOGENETIC RELATIONSHIPS clade, which, in turn, was joined by the clade A maximum parsimony analysis of the data matrix [(Trophonella + Boreotrophon) Paziella], the MP shown in Table 1 [PAUP* 4.0 beta version 10 Heuristic analysis groups the genera Haustrum, Trophonella, search, Accelerated transformation, TBR Swapping Boreotrophon,andPaziella into an unresolved polyt- algorithm] produced 475,972 equally parsimonious omy that is sister to Trophon and the polytomy trees (length 5 60; consistency index 5 0.783; retention containing Ocenebra. index 5 0.903]. A strict consensus of these trees is These results indicate that the type species of

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Trophonella is more closely related to Boreotrophon LITERATURE CITED than to the type species of Trophon and thus sup- BEU, A. G. 2004. Marine of oxygen isotope stages of port the segregation of Trophonella from Trophon the last 2 million years in New Zealand. Part 1: Revised at the generic level. They also confirm the conten- generic positions and recognition of warm-water and tions by various authors that Trophoninae, as cool-water migrants. Journal of the Royal Society of New defined by Cossmann (1903:9), is not monophyletic. Zealand 34(2):111–265. Based on the relationships of the type species of their CARCELLES, A. 1953. Cata´logo de la malacofauna antartica argentina. Anales del Museo Nahuel Huapi 3:155–250. respective type genera, either Trophoninae is the CASTELLANOS, Z. J. A. D. & N. LANDONI. 1993. Cata´logo sister taxon of Ocenebrinae or both are part of a larger descriptivo de la malacofauna marina magalla´nica 9. clade. Neogastropoda: Muricidae y Thaisidae. Comisio´n de In his phylogenetic study, Kool (1993b:fig. 30) Investigacio´nes Cientificas de la Provincia de Buenos considered Ocenebrinae to include Haustrum.Tan Aires. 26 pp. (2003) subsequently erected the subfamily Haustrinae COSSMANN, M. 1903. Muricidæ. Essais de Pale´onconchologie Compare´e 5:7–66, pls. 1–3. (with Haustrum as the type genus) to encompass a clade DELL, R. K. 1990. Antarctic Mollusca, with special reference of nonrapanine muricids endemic to Australia and to the Fauna of the Ross Sea. The Royal Soceity of New New Zealand. However, the relationship of this clade Zealand, Bulletin 27, iv + 311 pp. to Ocenebrinae and Rapaninae varied significantly, HAIN, S. G. 1990. Beitra¨ge zur Biologie der beschalten depending on the choice of outgroup. Beu (2004:216) Mollusken (Kl. Gastropoda und Bivalvia) des Weddell- concluded that it seemed preferable to retain the taxa meeres, Antarktis. Berichte zur Polarforschung 70:1–181. HAIN, S. & P. M. ARNAUD. 1992. Notes on the reproduction included by Tan in Haustrinae within the subfamily of high-Antarctic molluscs from the Weddell Sea. Polar Ocenebrinae. This broader delineation of Ocenebrinae Biology 12(2):303–312. includes the type species of Trophoninae in both HARASEWYCH, M. G. 1984. Comparative anatomy of four MP and NJ trees. The genera Trophonella and primitive muricacean gastropods: Implications for troph- Boreotrophon emerged as part of an unresolved onine phylogeny. American Malacological Bulletin 3(1): polytomy with Haustrum in the strict-consensus MP 11–26. HOUART, R. 1997. Description of Trophon eversoni n. sp. tree, but were depicted as being more distantly related (Gastropoda: Muricidae), a large trophonine from the in the NJ tree. Kerguelen Islands. Venus 56(1):9–13. Both Ocenebrinae and Trophoninae were proposed KOOL, S. P. 1987. Significance of Radular Characters in in the same publication (Cossmann, 1903:10), so Reconstruction of Thaidid Phylogeny (Neogastropoda: neither has priority. Although it is likely that these Muricacea). The Nautilus 101(3):117–132. two subfamilies will be synonymized, based on the KOOL, S. P. 1993a. The systematic position of the genus Nucella (Prosobranchia: Muricidae: Ocenebrinae). The close relationship between the type species of their type Nautilus 107(2):43–57. genera, a better resolved phylogeny, including a more KOOL, S. P. 1993b. Phylogenetic analysis of the Rapaninae comprehensive sampling of the more than 20 genus- (Neogastropoda: Muricidae). Malacologia 35(2):155–259. level taxa attributed to each of these two subfamilies, NUMANAMI, H. 1996. Taxonomic study on Antarctic gastro- will be required to delineate more precisely the pods collected by Japanese Antarctic research expeditions. membership of the clade and to identify its diagnostic Memoirs of National Institute of Polar Research, Series E synapomorphies. (Biology and Medical Science) No. 39, iv + 244 pp. OKUTANI, T. 1986. A note on Antarctic benthic mollusks collected with a beam-trawl from Breid Bay by the 25th Acknowledgments. We thank the following curators and Japanese Research Antarctic Expedition. Memoirs of collection managers for access to specimens in their collec- National Institute of Polar Research, Special Issue, tions: K. Way (NHM); A. Ware´n (NHRM); P. Bouchet No. 40, Pp. 277–287. and V. Heros (MNHN); P. Mikelsen (AMNH); B. Hausdorf PASTORINO, G. 2002. Systematics and phylogeny of the genus (ZMH), A. Tablado (MACN); C. Ituarte; (MLP) and R. Trophon Montfort, 1810 (Gastropoda: Muricidae) from Janssen (SM); P. Arnaud, Endoume, France. This work was Patagonia and Antarctica; morphological patterns. Bol- supported by a grant from the Consejo Nacional de lettino Malacologico, Supplement 4:127–134. Investigaciones Cientı´ficas y Te´cnicas (CONICET), Argen- POWELL, A. W. B. 1951. Antarctic and subAntarctic tina, which enabled the junior author to work in the Divi- Mollusca: Pelecypoda and Gastropoda. Discovery Re- sion of Mollusks, United States National Museum of ports 26:47–196. Natural History, Smithsonian Institution. Additional sup- POWELL, A. W. B. 1957. Mollusca of Kerguelen and port was provided in part by a Research Award from the Macquarie Islands. Reports of B.A.N.Z. Antarctic NSF-USAP United States Antarctic Program Grant Research Expedition, 1929–1931. Series B 6(7):107–150. [ANTO636408] and a grant-in-aid from the Conchologists POWELL, A. W. B. 1958. Mollusca from the Victoria-Ross of America and the Walter E. Sage Memorial Award. We Quadrants of Antarctica. Reports of B.A.N.Z. Antarctic are grateful to Dr. Yu. Kantor for his many helpful and Research Expedition, 1929–1931. Series B 6(9):167–215. insightful comments during the course of this research, and to POWELL, W. B. 1960. Antarctic and subantarctic mollusca. Roland Houart and Dr. Geerat Vermeij for their helpful Records of the Auckland Institute and Museum 5:117–193. reviews. RADWIN, G. E. & A. D’ATTILIO. 1976. Murex Shells of the

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World, An Illustrated Guide to the Muricidae. Stanford Parsimony (*and Other Methods). Version 4.0 Sinauer University Press, Stanford, California; x + 284, 32 pls pp. Associates, Sunderland, Massachusetts. SOLEM, A. 1972. Malacological application of Scanning TAN, K. S. 2003. Phylogenetic analysis and of some Electron Microscopy, II. Radular structure and function- southern Australian and New Zealand Muricidae (Mol- ing. The Veliger 14:327–336. lusca: Neogastropoda). Journal of Natural History 37: SWOFFORD, D. L. 2002. PAUP*. Phylogenetic Analysis Using 911–1028.

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